Behaviors can facilitate colonization of a novel environment, but the mechanisms underlying this process are poorly understood. On one hand, behavioral flexibility allows for an immediate response of colonizers to novel environments, which is critical to population establishment and persistence. On the other hand, integrated sets of behaviors that display limited flexibility can enhance invasion success by coupling behaviors with dispersal strategies that are especially important during natural range expansions. Direct observations of colonization events are required to determine the mechanisms underlying changes in behavior associated with colonization, but such observations are rare. Here, we studied changes in aggression on a large temporal and spatial scale across populations of two sister taxa of bluebirds (Sialia) to show that coupling of aggression and dispersal strongly facilitated the range expansion of western bluebirds across the northwestern United States over the last 30 years. We show that biased dispersal of highly aggressive males to the invasion front allowed western bluebirds to displace less aggressive mountain bluebirds. However, once mountain bluebirds were excluded, aggression of western bluebirds decreased rapidly across consecutive generations in concordance with local selection on highly heritable aggressive behavior. Further, the observed adaptive microevolution of aggression was accelerated by the link between dispersal propensity and aggression. Importantly, our results show that behavioral changes among populations were not caused by behavioral flexibility and instead strongly implicate adaptive integration of dispersal and aggression in facilitating the ongoing and rapid reciprocal range change of these species in North America.genetics of behavior ͉ geographical range ͉ competition ͉ colonization ͉ behavioral integration T he successful colonization of a novel environment is a crucial first step in many models of speciation, is central to source-sink population dynamics, and is a key component of natural and human-induced changes in species distributions. Yet, the mechanisms underlying the colonization of new environments, from the initial arrival of new individuals to population establishment, remain poorly understood (1-3). Because the success of colonizers depends on their ability to survive and reproduce in novel ecological conditions, they must either be preadapted to the new environment or be flexible enough to respond rapidly and adaptively to novel conditions (4).Behavioral flexibility is thought to aid colonization success by facilitating a rapid response to new conditions (5), thus enabling initial population persistence before adaptive evolution can occur (4, 6-8). In support of this idea, behavioral flexibility has been linked to the success of human-introduced invasive species (9, 10), yet its role in a natural range expansion has never been documented directly. Natural range expansion differs from colonization by invasive species, because, unlike invasive species that...

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As is the case with any metaphor, parental effects mean different things to different biologists—from developmental induction of novel phenotypic variation to an evolved adaptation, and from epigenetic transference of essential developmental resources to a stage of inheritance and ecological succession. Such a diversity of perspectives illustrates the composite nature of parental effects that, depending on the stage of their expression and whether they are considered a pattern or a process, combine the elements of developmental induction, homeostasis, natural selection, epigenetic inheritance and historical persistence. Here, we suggest that by emphasizing the complexity of causes and influences in developmental systems and by making explicit the links between development, natural selection and inheritance, the study of parental effects enables deeper understanding of developmental dynamics of life cycles and provides a unique opportunity to explicitly integrate development and evolution. We highlight these perspectives by placing parental effects in a wider evolutionary framework and suggest that far from being only an evolved static outcome of natural selection, a distinct channel of transmission between parents and offspring, or a statistical abstraction, parental effects on development enable evolution by natural selection by reliably transferring developmental resources needed to reconstruct, maintain and modify genetically inherited components of the phenotype. The view of parental effects as an essential and dynamic part of an evolutionary continuum unifies mechanisms behind the origination, modification and historical persistence of organismal form and function, and thus brings us closer to a more realistic understanding of life's complexity and diversity.

Extreme environments are closely associated with phenotypic evolution, yet the mechanisms behind this relationship are poorly understood. Several themes and approaches in recent studies significantly further our understanding of the importance that stress-induced variation plays in evolution. First, stressful environments modify (and often reduce) the integration of neuroendocrinological, morphological and behavioural regulatory systems. Second, such reduced integration and subsequent accommodation of stress-induced variation by developmental systems enables organismal 'memory' of a stressful event as well as phenotypic and genetic assimilation of the response to a stressor. Third, in complex functional systems, a stress-induced increase in phenotypic and genetic variance is often directional, channelled by existing ontogenetic pathways. This accounts for similarity among individuals in stress-induced changes and thus significantly facilitates the rate of adaptive evolution. Fourth, accumulation of phenotypically neutral genetic variation might be a common property of locally adapted and complex organismal systems, and extreme environments facilitate the phenotypic expression of this variance. Finally, stress-induced effects and stress-resistance strategies often persist for several generations through maternal, ecological and cultural inheritance. These transgenerational effects, along with both the complexity of developmental systems and stressor recurrence, might facilitate genetic assimilation of stress-induced effects. Accumulation of phenotypically neutral genetic variance by developmental systems and phenotypic accommodation of stress-induced effects, together with the inheritance of stress-induced modifications, ensure the evolutionary persistence of stress-response strategies and provide a link between individual adaptability and evolutionary adaptation.

▪ Abstract  The extent and diversity of sexual dichromatism in birds is thought to be due to the intensity of current sexual selection on the plumage ornamentation of males and females. This view leads to an expectation of concordance between ecological conditions and sexual dichromatism. Yet, because expression of dichromatism is the result of not only current selection, but also historical patterns of development, function, and selection, the concordance between ecology and current sexual dichromatism is not straightforward. Recent studies have revealed a number of trends in the evolution of avian sexual ornamentation that seem contrary to what is expected if current sexual selection is the primary force shaping dichromatism. For example, change in sexual dichromatism is often the result of evolutionary changes in female rather than male ornamentation. Moreover, sexual dichromatism is often an ancestral rather than a derived state; current expression of dichromatism is frequently the result of selection for lesser ornamentation in one sex and not for ornament elaboration. Loss and gain of sexual ornamentation sometimes precedes changes in preference for sexual ornamentation, and sexual ornaments can have high evolutionary lability despite their developmental and functional complexity. These findings emphasize that phylogenetic reconstructions must play a central role in attempts to understand the function and evolution of sexual dichromatism. With a historical perspective, one can test the relative importance of direct selection, indirect selection, and drift in relation to changes of sexual dichromatism. If sexual selection is invoked, the mechanisms of sexual selection can be explored by examining the concordance between the elaboration of ornamentation and the preferences for ornamentation across species and by tracing phylogenetic trajectories of sexual ornaments. Finally, placing physiological, genetic, and developmental mechanisms of sexual ornamentation into such a phylogenetic framework will enable greater inference about the past evolution and current function of sexual dichromatism in birds.

Most species of birds can lay only one egg per day until a clutch is complete, and the order in which eggs are laid often has strong and sex-specific effects on offspring growth and survival. In two recently established populations of the house finch (Carpodacus mexicanus) in Montana and Alabama, breeding females simultaneously adjusted the sex and growth of offspring in relation to their position in the laying order, thereby reducing the mortality of sons and daughters by 10 to 20% in both environments. We show experimentally that the reduction in mortality is produced by persistent and sex-specific maternal effects on the growth and morphology of offspring. These strong parental effects may have facilitated the rapid adaptive divergence among populations of house finches.

The social environment is a critical determinant of fitness and, in many taxa, is shaped by an individual's behavioral discrimination among social contexts, suggesting that animals can actively influence the selection they experience. In competition to attract females, males may modify sexual selection by choosing social environments in which they are more attractive relative to rivals. Across the population, such behaviors should influence sexual selection patterns by altering the relationship between male mating success and sexual ornament elaboration. Here we use network analysis to examine patterns of male social behavior in relation to plumage ornamentation and mating success in a free-living population of house finches. During the nonbreeding season, less elaborate males changed associations with distinct social groups more frequently, compared to more elaborate males that showed greater fidelity to a single social group. By the onset of pair formation, socially labile males effectively increased their attractiveness relative to other males in the same flocks. Consequently, males that frequently moved between social groups had greater pairing success than less social individuals with equivalent sexual ornamentation. We discuss these results in relation to conditional mating tactics and the role of social behavior in evolutionary change by sexual selection.

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